Allele Dmel\ato¹

General Information
Symbol	Dmel\ato¹	Species	D. melanogaster
Name		FlyBase ID	FBal0034197
Feature type	allele	Associated gene	Dmel\ato
Also Known As	ato1

Map ( GBrowse )
Allele class	loss of function allele, amorphic allele - genetic evidence
Mutagen	ethyl methanesulfonate
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
Update Feed	Click the icon below to subscribe to this FlyBase record and receive updates automatically through your feed reader.
FB2013_03
FB2013_02
All updates	Click here to see a list of all updates to this record from FB2010_08 and on.
Nature of the Allele
Allele class	amorphic allele - genetic evidence (Jarman et al., 1994, Brown et al., 1996, Gupta and Rodrigues, 1997) loss of function allele (Jarman et al., 1995, Reddy et al., 1997, zur Lage et al., 1997)
Mutagen	ethyl methanesulfonate (Jarman et al., 1994)
Mutations Mapped to the Genome

Type Location Additional Notes References point mutation 3R:4,103,920..4,103,920 evidence=experimental na_change=G4103920A pr_change=A25T\|ato-PA reported_pr_change=A25T comment=One of three predicted amino acid changes in the mutant relative to the parent chromosome. comment=Site of nucleotide substitution in mutant inferred by FlyBase based on reported amino acid change. (Jarman et al., 1994) point mutation 3R:4,104,606..4,104,606 evidence=experimental na_change=A4104606Y pr_change=K253N\|ato-PA reported_pr_change=K253N comment=One of three predicted amino acid changes in the mutant relative to the parent chromosome. comment=Site of nucleotide substitution in mutant inferred by FlyBase based on reported amino acid change. (Jarman et al., 1994) point mutation 3R:4,104,629..4,104,629 evidence=experimental na_change=A4104629T pr_change=N261I\|ato-PA reported_pr_change=N261I comment=One of three predicted amino acid changes in the mutant relative to the parent chromosome. comment=Site of nucleotide substitution in mutant inferred by FlyBase based on reported amino acid change. (Jarman et al., 1994)
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference Deficient in the DNA binding domain. (Dokucu et al., 1996) Amino acid replacement: N261I. Amino acid replacement: K253N. Amino acid replacement: A25T. Mutations are in a motif of the basic domain (N261I), a short distance proximally on the edge of the basic domain (K253N) and near the N terminus respectively (A25T). (Jarman et al., 1994)
Cytology
Phenotypic Data
Phenotypic Class
	auditory perception defective (Eberl et al., 2000, Cosetti et al., 2008) behavior defective \| larval stage (Wu et al., 2011) behavior defective \| recessive (Jarman et al., 1995) locomotor behavior defective (with Df(3R)p13) (Caldwell et al., 2003) neuroanatomy defective (Hassan et al., 2000) neuroanatomy defective (with Df(3R)p13) (Jhaveri and Rodrigues, 2002) neuroanatomy defective \| embryonic stage (Grillenzoni et al., 2007) neurophysiology defective (Eberl et al., 2000, Cosetti et al., 2008) partially lethal - majority die \| recessive (Jarman et al., 1994) song defective (Tauber and Eberl, 2001) touch sensitivity defective (with Df(3R)p13) (Caldwell et al., 2003) uncoordinated \| recessive (Ma and Jarman, 2011) visible (Sun et al., 1998) visible (with ato²) (White and Jarman, 2000) visible (with Df(3R)p13) (Siddall et al., 2003, Reddy et al., 1997) visible \| recessive (Goulding et al., 2000, White and Jarman, 2000, Gupta and Rodrigues, 1997, Chanut et al., 2002, Jarman et al., 1994) visual behavior defective (Jarman et al., 1994)
Phenotype Manifest In
	abdominal 1 lateral pentascolopidial chordotonal organ lch5 (Chien et al., 1996) abdominal 2 lateral pentascolopidial chordotonal organ lch5 (Chien et al., 1996) abdominal 3 lateral pentascolopidial chordotonal organ lch5 (Chien et al., 1996) abdominal 4 lateral pentascolopidial chordotonal organ lch5 (Chien et al., 1996) abdominal 5 lateral pentascolopidial chordotonal organ lch5 (Chien et al., 1996) abdominal 6 lateral pentascolopidial chordotonal organ lch5 (Chien et al., 1996) abdominal 7 lateral pentascolopidial chordotonal organ lch5 (Chien et al., 1996) abdominal lateral pentascolopidial chordotonal organ lch5 (Elstob et al., 2001) abdominal lateral pentascolopidial chordotonal organ lch5 (with Df(3R)p13) (Elstob et al., 2001) adult antennal nerve (Gupta and Rodrigues, 1997, Eberl et al., 2000) adult brain (Hassan et al., 2000) antenna (Dong et al., 2002) antenna (with Df(3R)p13) (Jhaveri and Rodrigues, 2002) antenna & neuron (Sen et al., 2004) antennal lobe (with Df(3R)p13) (Jhaveri and Rodrigues, 2002) antennal lobe glomerulus (with Df(3R)p13) (Jhaveri et al., 2000, Jhaveri and Rodrigues, 2002) antennal segment 3 & fascicle (with Df(3R)p13) (Jhaveri et al., 2000) basiconic sensillum of sacculus (Gupta and Rodrigues, 1997) Bolwig organ (Daniel et al., 1999) C1 chordotonal organ precursor cell (with Df(3R)p13) (Brodu et al., 2004) chordotonal organ (Jarman et al., 1995, zur Lage et al., 1997) chordotonal organ \| embryonic stage (Ben-Arie et al., 2000) embryonic/larval brain (Hassan et al., 2000) embryonic/larval oenocyte (Rusten et al., 2001, Elstob et al., 2001) embryonic/larval oenocyte (with Df(3R)p13) (Elstob et al., 2001, Brodu et al., 2002) embryonic/larval oenocyte precursor (Witt et al., 2010) embryonic/larval oenocyte precursor (with Df(3R)p13) (Brodu et al., 2004) embryonic antennal sense organ (Grillenzoni et al., 2007) eye (White and Jarman, 2000, Sun et al., 1998, Chanut et al., 2002, Sun et al., 2000, Jarman et al., 1994) eye (with ato²) (White and Jarman, 2000) eye (with ato¹⁰⁹⁰) (Melicharek et al., 2008) eye (with Df(3R)p13) (Siddall et al., 2003, Yu et al., 2012) eye \| heat sensitive (with ato¹⁰⁹⁰) (Distefano et al., 2012) eye-antennal disc (Jarman et al., 1995, Jarman et al., 1994) eye disc (Lim and Choi, 2004, Quan et al., 2004) eye photoreceptor cell (with ato²) (White and Jarman, 2000) eye photoreceptor cell & axon (with ato²) (White and Jarman, 2000) glial cell (with Df(3R)p13) (Sen et al., 2004) glial cell & antenna (with Df(3R)p13) (Jhaveri and Rodrigues, 2002) glial cell & antennal lobe (with Df(3R)p13) (Jhaveri and Rodrigues, 2002) interneuron & antennal lobe (with Df(3R)p13) (Jhaveri and Rodrigues, 2002) interommatidial bristle (Jarman et al., 1994) Johnston organ (Dong et al., 2002, Jarman et al., 1995, Eberl et al., 2000) Johnston organ (with Df(3R)p13) (Senthilan et al., 2012) maxillary palp sensillum basiconicum (Gupta and Rodrigues, 1997) morphogenetic furrow (Jarman et al., 1995) morphogenetic furrow & filopodium \| somatic clone (Chou and Chien, 2002) morphogenetic furrow \| somatic clone (Chen and Chien, 1999) neuron of aristal sensillum (with Df(3R)p13) (Jhaveri et al., 2000) ocellus (Jarman et al., 1994) oenocyte primordium (with Df(3R)p13) (Brodu et al., 2004) olfactory neuron & embryonic antennal sense organ (Grillenzoni et al., 2007) ommatidial precursor cluster \| somatic clone (Brown et al., 2006) ommatidium (Sun et al., 1998, Sun et al., 2000) ommatidium (with ato²) (White and Jarman, 2000) ommatidium (with Df(3R)p13) (Yu et al., 2012) ommatidium \| somatic clone (Chou and Chien, 2002) optic lobe (Jarman et al., 1994) photoreceptor cell (Jarman et al., 1995, Nolo et al., 2000, Jarman et al., 1994) photoreceptor cell R8 (White and Jarman, 2000, Sun et al., 2003) pioneer neuron & adult antennal nerve & pupa (with Df(3R)p13) (Jhaveri et al., 2004) sacculus (Gupta and Rodrigues, 1997) sacculus (with Df(3R)p13) (Reddy et al., 1997) scolopidial neuron (Jarman et al., 1995, Huang et al., 2000) sensillum coeloconicum (Goulding et al., 2000) sensillum coeloconicum of antennal segment 3 (Gupta and Rodrigues, 1997) sensillum coeloconicum of antennal segment 3 (with ato²) (Jhaveri and Rodrigues, 2002) sensillum coeloconicum of antennal segment 3 (with Df(3R)p13) (Jhaveri and Rodrigues, 2002, Reddy et al., 1997) sensory mother cell (Rusten et al., 2001) sensory mother cell & antennal disc (zur Lage et al., 2003) third segment of antenna (Gopfert et al., 2002) third segment of antenna (with Df(3R)p13) (Gopfert et al., 2002) wing (Tauber and Eberl, 2001)
Detailed Description
	Statement Reference ato[1]/ato[1090] mutants exhibit a small, rough eye phenotype at 25[o]C. (Distefano et al., 2012) ato[1]/Df(3R)p13 flies lack the Johnston's organ. (Senthilan et al., 2012) Hemizygous ato[1]/Df(3R)p13 escapers are nearly or completely eyeless, displaying only a few ommatidia when eye tissue is present. (Yu et al., 2012) Adult ato[1] flies show uncoordinated behaviour. They have a dramatically reduced climbing ability in bang tests compared with wild type controls. (Ma and Jarman, 2011) Mutant larvae do not show a normal response to vibration, showing a small decrease in crawling speed with no head turning. (Wu et al., 2011) The total number of oenocytes that form is significantly decreased in ato[1] mutant embryos as compared to controls. Control embryos have clusters of approximately six oenocytes in each of seven abdominal segments. In contrast, many segments of ato[1] mutant embryos completely lack oenocytes. When oenocytes do develop in ato[1] mutants, fewer of them form than in control segments. (Witt et al., 2010) 90.5% of abdominal segments contain an lch5 neuron in ato[1] homozygous embryos. (zur Lage and Jarman, 2010) Mutants show no response to an auditory stimulus mimicking flies' courtship song. (Cosetti et al., 2008) Transheterozygous ato[1090]/ato[1] mutants produce small rough eyes at 25[o]C. (Melicharek et al., 2008) The number of neurons in the dorsal organ ganglion are reduced in mutant embryos compared to wild type. The number of olfactory receptor neurons in the DO ganglion is reduced in the mutant embryos (average number is 8 compared to the wild-type number of 21). (Grillenzoni et al., 2007) ato¹ Minute clones in the eye disc fail to form the arcs and rosettes of cells in the morphogenetic furrow that are seen in wild-type eye discs. (Brown et al., 2006) ato[1]/Df(3R)p13 embryos lack C1 chordotonal organ precursor cells and have no larval oenocyte precursors. (Brodu et al., 2004) In the adult olfactory system, ato specifies a subset of neurons that are the first to develop and appear to guide the rest of the axons into the lobe. In ato¹/Df(3R)p13 animals, "pioneer" antennal sensory neurons fail to form and the remaining antennal sensory neurons stall at the entry to the olfactory lobe. (Jhaveri et al., 2004) Morphogenetic furrow formation can occur in mutant eye discs and the furrow progresses anteriorly to a certain distance, although retinal differentiation fails to occur. (Lim and Choi, 2004) ato¹/Df(3R)p13 mutant animals lack a large fraction of glia. (Sen et al., 2004) ato¹/Df(3R)p13 larvae have reduced touch sensitivity compared to wild-type. Locomotion in these larvae is aberrant: compared to controls they are slower and they pause, turn and retreat more often. These defects in locomotion arise at least partially from defective peristaltic motion: maximum larval length, stride period and % positive and negative flow are all defective. (Caldwell et al., 2003) ato¹/Df(3R)p13 flies have an eye that is a patch of pigment with few sensory hairs. (Siddall et al., 2003) In the pupal antennal imaginal disc of mutant animals, the first and second waves of sensory organ precursor formation do not occur. (zur Lage et al., 2003) ato¹/Df(3R)p13 embryos lack oenocytes. (Brodu et al., 2002) The eyes of homozygotes are reduced to a slit of pigment cells. (Chanut et al., 2002) Ommatidial clusters situated posterior to ato¹ clones display over-rotation phenotypes at a high frequency. Cytoplasmic extensions, extending from the morphogenetic furrow into these clones, follow convoluted paths rather than being straight and aligned with each other as in wild-type. (Chou and Chien, 2002) ato¹ mutants lack the Johnston's organ and the circular outline of the joint between antennal segments 2 and 3. (Dong et al., 2002) The mechanical response of the antenna to sound is severely affected in homozygous and ato[1]/Df(3R)p13 flies compared to wild-type controls. The response of the entire antenna to sound in homozygous and ato[1]/Df(3R)p13 flies is similar to that observed on the head capsule with respect to both its amplitude and frequency composition. Neither the funiculus nor the arista vibrate in a coherent manner with sound (in contrast to wild type), demonstrating that the entire antenna is immobile and stiff in the mutant animals. Homozygous and ato[1]/Df(3R)p13 flies show anatomical defects in the antennal joints. The hook that is normally present at the proximal end of the funicular stalk is missing in the mutant animals, and the funicular stalk enters the pit of the pedicel only in part, connecting broadly to both the inner wall and the anterior edge of the pedicel. The V-shaped cuticular rims that are present on either side of the hook in cross-sections at the connection between the pedicel and funiculus in wild-type animals are missing in the mutant flies, and the thin joint membranes that surround the hook and the rims in wild-type flies are missing, with the intersegmental cuticle being constantly thick in the mutant flies. (Gopfert et al., 2002) Antennal nerves stall at the periphery of the olfactory lobe up until the mid-pupal stage in ato¹/Df(3R)p13 animals, with only very few fibres projecting into the lobe. The antennal nerves do enter the olfactory lobe later in development, but they remain disorganised and no obvious glomerular structures are seen. The antennal commissure fails to form. Olfactory lobe associated interneurons are normal. The distribution of glial cells within the antennal lobe is normal, but the glial processes show excessive branching. The number of peripheral glia associated with the olfactory neurons in the antenna is reduced in ato¹/Df(3R)p13 pupae and the 3 fascicle that are normally seen in wild-type animals are merged into a single bundle. Projection interneurons are present in the antennal lobes of ato¹/Df(3R)p13 adults, but they are unpatterned, in contrast to wild type. ato¹/Df(3R)p13 antenna completely lack coeloconic sensilla. ato¹/ato² antenna have a small number of coeloconic sensilla. The antennal lobes have normal glomeruli and the inter-antennal commissure is present. The elaboration of glial cell processes within the antennal lobe is normal. (Jhaveri and Rodrigues, 2002) ato¹/Df(3R)p13 embryos have hemisegments in which the lch5 array is either completely missing or reduced to a single lateral chordotonal organ. Oenocyte formation is completely abolished in those segments lacking all lateral chordotonal organs. In the majority of cases where a single lateral chordotonal organ is formed, it is associated with an oenocyte cluster of normal or somewhat reduced cell number. Similar phenotypes are seen in ato¹ homozygotes, although the phenotype is less penetrant. (Elstob et al., 2001) Oenocytes do not form in homozygous embryos, where the formation of the primary sensory organ precursors (SOPs) is compromised in most segments. In segments where remnant SOPs do develop, oenocytes can develop. (Rusten et al., 2001) Mutant males demonstrate vigorous courtship including courtship songs. 66% of mutant males use both of their wings simultaneously during courtship song production (in contrast to wild-type males which vibrate only one wing at a time). Relative amplitude of the sine song and pulse number is higher than normal in mutant males. Pulse duration is significantly longer than wild-type. (Tauber and Eberl, 2001) Mutant embryos lack all chordotonal neurons. (Ben-Arie et al., 2000) The second antennal segment is devoid of scolopidia. The antennal nerve is present. No sound evoked potential can be recorded from the antennal nerve. (Eberl et al., 2000) The dorsal, ventral brain and ventral lobular clusters (that normally express ato) are present in homozygous and hemizygous brains. In third larval instar brains there are severe defects in dorsal cluster (DC) position and organisation, and the cells appear to be less tightly clustered then normal. The axons are clearly defasciculated. These defects are seen with about 15% penetrance. The DC axons form a commissural tract. In adult brains, in addition to the positioning defects of the DC, the arborisation pattern of the DC over the lobula is severely impaired in hemizygous flies. Most axons enter the lobula either ventrally or dorsally and show very limited branching, failing to form a proper "fan-shaped" pattern. (Hassan et al., 2000) ato¹/Df(3R)p13 animals show a striking change in fascicular pattern of sensory neurons and glia in the third antennal segment. While all three branches are identifiable, they exit the antenna in a single bundle rather than distinct fascicles. The aristal neurons also fail to form. The number of glial cells present along the exiting sensory projections is also significantly reduced. ato¹/Df(3R)p13 animals also show a dramatic disorganisation of the antennal glomeruli. In most antennal lobes examined, no glomeruli can be discerned at all. In a few cases one or two glomeruli are apparent. (Jhaveri et al., 2000) Photoreceptor development is aberrant in ato¹ eye-antennal discs. (Nolo et al., 2000) The eyes of ato¹ mutant flies are totally devoid of ommatidia. (Sun et al., 2000) ato¹/ato² flies have severely reduced eyes, which nevertheless contain ommatidia (approximately 200-300 in males, 5-15 in females). The ommatidia mostly contain fewer photoreceptors than wild type. The photoreceptor clusters are variably reduced, containing from 1-8 photoreceptors and any or all of the recruited photoreceptors can be affected. Photoreceptor axons navigate to the optic stalk poorly. This axon pathfinding defect is more severe in females. Those axons that reach the optic stalk are able to continue to their target sites within the optic lobe. Homozygous flies lack the compound eye, since the ommatidial founder cells (R8 photoreceptors) are not selected in these animals. (White and Jarman, 2000) Progression of the morphogenetic furrow is slightly delayed across homozygous clones in the eye. (Chen and Chien, 1999) Mutant embryos lack the Bolwig's organ. (Daniel et al., 1999) No ommatidia are formed in the presumptive eye field of homozygous flies, except for a few pigment cells and bristles. Flies carrying one or two copies of ato^t5'eye show no rescue of the ato¹ eye phenotype. Flies carrying one or two copies of both ato^t5'eye and ato^t3'F.5.8 show rescue of the ato¹ eye phenotype; flies carrying one copy of each construct typically have eyes containing 40% of the normal number of ommatidia, while flies carrying two copies of each construct have almost wild-type eyes. (Sun et al., 1998) Hemizygous antenna lack all the sensilla coeloconica. Maxillary palps lack sensilla basiconica. Antenna sacculus is absent, instead a few sensilla basiconica are present. The axonal bundle 2 fails to emerge out of the third antennal segment. Loss of sc and ac does not affect the hemizygous phenotype. (Gupta and Rodrigues, 1997) ato¹/Df(3R)p13 flies completely lack sensilla coelonica on the surface of the third antennal segment, while the numbers and morphology of the sensilla basiconica and trichodea are normal. The sacculus is missing. (Reddy et al., 1997) In mutant embryos all chordotonal organs are absent except for one scolopidium of lch5. (zur Lage et al., 1997) The lateral five chordotonal organ (lch5) neurons are missing in each abdominal segment. Most of the lch5 neurons are restored in the thoracic segment T2 and the odd-numbered abdominal segments when ato^Scer\UAS.cCa is expressed in ato¹ flies using Scer\GAL4^h-1J3. ato::sc^{bHLH.ato.Scer\UAS} and ato::sc^{b.ato.Scer\UAS} restore many of the lch5 neurons in the odd-numbered abdominal segments A1 and A3 when expressed in ato¹ flies using Scer\GAL4^h-1J3. sc^Scer\UAS.cCa does not rescue the ato¹ mutant phenotype when expressed using Scer\GAL4^h-1J3. (Chien et al., 1996) Embryos lack almost all chordotonal organs of the thorax and abdomen. The remaining PNS is largely unaffected. Mutant larvae can hatch and survive to adulthood. Adult flies lack chordotonal organs associated with femur, wing base or ventral abdomen, and also lack Johnston's organ. Flies are clumsy on their feet and attempt to fly only with extreme reluctance. Chordotonal organ precursors are clearly and specifically missing in the leg, wing and antennal discs. No photoreceptors form, though a partial morphogenetic furrow remains in the mutant eye discs. Extensive cell death occurs in the larval eye disc posterior to the morphogenetic furrow. BrdU incorporation studies suggest that the second round or replication, which occurs in wild type posterior to the morphogenetic furrow, only occurs in the mutant as long as the furrow is moving. (Jarman et al., 1995) Homozygotes and hemizygotes are almost eyeless, compound eye is a small red patch with a smooth surface and a few ommatidial bristles, and also lack ocelli. Photoreceptors are completely absent and the optic lobe is reduced. Eye antennal disc from third instar larvae shows a remnant of the morphogenetic furrow and extensive cell death in regions posterior to this crease. X ray induced homozygous clones in developing eye discs of heterozygous larvae result in a scar due to absence of ommatidia. (Jarman et al., 1994)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Suppressed by
Statement Reference ato¹, ato^t3'F.5.8 has visible phenotype, suppressible \| partially by sc^t5'eye (Sun et al., 2000) ato¹, ato^t5'eye has visible phenotype, suppressible \| partially by sc^t3'F.5.8 (Sun et al., 2000) ato¹/Df(3R)p13 has visible phenotype, suppressible \| partially by BacA\p35^GMR.PH (Siddall et al., 2003) ato¹ has visible \| recessive phenotype, suppressible \| partially by amos^Roi-1/amos[+] (Chanut et al., 2002) ato¹ has visible phenotype, suppressible \| partially by sc^Scer\UAS.cCa/Scer\GAL4⁷ (Sun et al., 2000)
NOT suppressed by
Statement Reference ato¹ has visible phenotype, non-suppressible by sc^t3'F.5.8 (Sun et al., 2000) ato¹ has visible phenotype, non-suppressible by sc^t3'F.5.8/sc^t5'eye (Sun et al., 2000) ato¹ has visible phenotype, non-suppressible by sc^t5'eye (Sun et al., 2000)
Enhancer of
Statement Reference ato¹/ato[+] is an enhancer of visible phenotype of ro^DOM (Chanut et al., 2000)
Other
Statement Reference ato¹, cato⁵³⁶ has neuroanatomy defective \| embryonic stage phenotype (zur Lage and Jarman, 2010) ato¹, da⁹/da[+] has visible \| dominant phenotype (Gupta and Rodrigues, 1997) ato¹/ato[+], da⁹ has visible \| dominant phenotype (Gupta and Rodrigues, 1997) Scer\GAL4⁷, ato¹, sc^Scer\UAS.cCa has visible phenotype (Sun et al., 2000, Sun et al., 2000)
Phenotype Manifest In
Enhanced by
Statement Reference ato¹/ato¹⁰⁹⁰ has eye \| heat sensitive phenotype, enhanceable by lilli[+]/lilli^GD17 (Distefano et al., 2012) ato¹/ato¹⁰⁹⁰ has eye \| heat sensitive phenotype, enhanceable by lilli^AG5/lilli[+] (Distefano et al., 2012)
Suppressed by
Statement Reference ato¹, ato^t3'F.5.8 has ommatidium phenotype, suppressible \| partially by sc^t5'eye (Sun et al., 2000) ato¹, ato^t5'eye has ommatidium phenotype, suppressible \| partially by sc^t3'F.5.8 (Sun et al., 2000) ato¹, ato^t5'eye has photoreceptor cell R8 phenotype, suppressible \| partially by sc^t3'F.5.8 (Sun et al., 2000) ato¹/Df(3R)p13 has embryonic/larval oenocyte phenotype, suppressible by rho^{Scer\UAS.cdCa}/Scer\GAL4^en-e16E (Brodu et al., 2002) ato¹/Df(3R)p13 has embryonic/larval oenocyte precursor phenotype, suppressible by rho^{Scer\UAS.cdCa}/Scer\GAL4^en-e16E (Brodu et al., 2004) ato¹/Df(3R)p13 has eye phenotype, suppressible \| partially by BacA\p35^GMR.PH (Siddall et al., 2003) ato¹/Df(3R)p13 has eye phenotype, suppressible by ato::Bmor\ato^{BmDm.Scer\UAS}/Scer\GAL4^ato.eye (Yu et al., 2012) ato¹/Df(3R)p13 has eye phenotype, suppressible by ato::Bmor\ato^{DmBm.Scer\UAS}/Scer\GAL4^ato.eye (Yu et al., 2012) ato¹/Df(3R)p13 has eye phenotype, suppressible by Bmor\ato^Scer\UAS.cYb/Scer\GAL4^ato.eye (Yu et al., 2012) ato¹/Df(3R)p13 has oenocyte primordium phenotype, suppressible by rho^{Scer\UAS.cdCa}/Scer\GAL4^en-e16E (Brodu et al., 2004) ato¹ has chordotonal organ \| embryonic stage phenotype, suppressible \| partially by Mmus\Atoh1^Scer\UAS.cBa/Scer\GAL4^hs.PB (Ben-Arie et al., 2000) ato¹ has chordotonal organ phenotype, suppressible by Mmus\Atoh1^Scer\UAS.cBa/Scer\GAL4^ato.3.6 (Wang et al., 2002) ato¹ has eye phenotype, suppressible \| partially by amos^Roi-1/amos[+] (Chanut et al., 2002) ato¹ has eye phenotype, suppressible \| partially by sc^Scer\UAS.cCa/Scer\GAL4⁷ (Sun et al., 2000) ato¹ has ommatidial precursor cluster \| somatic clone phenotype, suppressible \| partially by Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF (Brown et al., 2006) ato¹ has ommatidial precursor cluster \| somatic clone phenotype, suppressible by Egfr::tor^act.Scer\UAS/Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF (Brown et al., 2006) ato¹ has ommatidium phenotype, suppressible \| partially by Mmus\Atoh7^Scer\UAS.cSa/Scer\GAL4⁷ (Sun et al., 2003) ato¹ has ommatidium phenotype, suppressible \| partially by sc^Scer\UAS.cCa/Scer\GAL4⁷ (Sun et al., 2000, Sun et al., 2003) ato¹ has ommatidium phenotype, suppressible \| partially by Xlae\ath5a^Scer\UAS.cSa/Scer\GAL4⁷ (Sun et al., 2003) ato¹ has ommatidium phenotype, suppressible \| partially by Xlae\neuroD^Scer\UAS.cSa/Scer\GAL4⁷ (Sun et al., 2003) ato¹ has photoreceptor cell R8 phenotype, suppressible \| partially by Xlae\ath5a^Scer\UAS.cSa/Scer\GAL4⁷ (Sun et al., 2003) ato¹ has photoreceptor cell R8 phenotype, suppressible \| partially by Xlae\neuroD^Scer\UAS.cSa/Scer\GAL4⁷ (Sun et al., 2003) ato¹ has scolopidial neuron phenotype, suppressible by amos^Scer\UAS.cHa/Scer\GAL4^h-1J3 (Huang et al., 2000)
NOT suppressed by
Statement Reference ato¹, ato^t3'F.5.8 has photoreceptor cell R8 phenotype, non-suppressible by sc^t5'eye (Sun et al., 2000) ato¹/Df(3R)p13 has ommatidium phenotype, non-suppressible by Bmor\ato^Scer\UAS.cYb/Scer\GAL4^ato.eye (Yu et al., 2012) ato¹ has eye disc phenotype, non-suppressible by Mmus\Neurog1^Scer\UAS.cQa/Scer\GAL4⁷ (Quan et al., 2004) ato¹ has interommatidial bristle phenotype, non-suppressible by Mmus\Atoh7^Scer\UAS.cSa/Scer\GAL4⁷ (Sun et al., 2003) ato¹ has ommatidium phenotype, non-suppressible by sc^t3'F.5.8 (Sun et al., 2000) ato¹ has ommatidium phenotype, non-suppressible by sc^t3'F.5.8/sc^t5'eye (Sun et al., 2000) ato¹ has ommatidium phenotype, non-suppressible by sc^t5'eye (Sun et al., 2000) ato¹ has photoreceptor cell R8 phenotype, non-suppressible by sc^Scer\UAS.cCa/Scer\GAL4⁷ (Sun et al., 2000) ato¹ has photoreceptor cell R8 phenotype, non-suppressible by sc^t3'F.5.8 (Sun et al., 2000) ato¹ has photoreceptor cell R8 phenotype, non-suppressible by sc^t3'F.5.8/sc^t5'eye (Sun et al., 2000) ato¹ has photoreceptor cell R8 phenotype, non-suppressible by sc^t5'eye (Sun et al., 2000)
Enhancer of
Statement Reference ato¹/ato[+] is an enhancer of eye phenotype of ro^DOM (Chanut et al., 2000)
NOT Enhancer of
Statement Reference ato¹/ato[+] is a non-enhancer of ommatidium phenotype of amos^Roi-1 (Chanut et al., 2002)
Suppressor of
Statement Reference ato¹/Df(3R)p13 is a suppressor \| partially of glial cell \| ectopic & antennal disc phenotype of BacA\p35^Scer\UAS.cHa, Scer\GAL4^Dll-981 (Sen et al., 2004)
NOT Suppressor of
Statement Reference ato¹/ato[+] is a non-suppressor of ommatidium phenotype of amos^Roi-1 (Chanut et al., 2002) ato¹/Df(3R)p13 is a non-suppressor of embryonic/larval oenocyte \| supernumerary phenotype of Scer\GAL4^en-e16E, rho^{Scer\UAS.cdCa} (Brodu et al., 2004) ato¹/Df(3R)p13 is a non-suppressor of embryonic/larval oenocyte precursor \| supernumerary phenotype of Scer\GAL4^en-e16E, rho^{Scer\UAS.cdCa} (Brodu et al., 2004) ato¹/Df(3R)p13 is a non-suppressor of oenocyte primordium phenotype of Scer\GAL4^en-e16E, rho^{Scer\UAS.cdCa} (Brodu et al., 2004)
Other
Statement Reference amos¹, ato¹ has sensory neuron \| somatic clone phenotype (zur Lage et al., 2003, zur Lage et al., 2003, zur Lage et al., 2003) ato¹, cato⁵³⁶ has lch5 neuron phenotype (zur Lage and Jarman, 2010) ato¹, da[+]/da¹⁰ has sensillum coeloconicum of antennal segment 3 phenotype (Goulding et al., 2000) ato¹, da⁹/da[+] has sensillum coeloconicum of antennal segment 3 phenotype (Gupta and Rodrigues, 1997) ato¹, da¹⁰ has sensillum coeloconicum of antennal segment 3 phenotype (Goulding et al., 2000) ato¹/ato[+], da⁹ has sensillum coeloconicum of antennal segment 3 phenotype (Gupta and Rodrigues, 1997) Df(1)sc-B57, ato¹ has dorsal multidendritic neuron phenotype (Huang et al., 2000) Df(2L)GR4, ato¹ has peripheral nervous system phenotype (Brewster et al., 2001) Scer\GAL4⁷, ato¹, sc^Scer\UAS.cCa has interommatidial bristle phenotype (Sun et al., 2003) Scer\GAL4⁷, ato¹, sc^Scer\UAS.cCa has macrochaeta \| ectopic phenotype (Sun et al., 2000, Sun et al., 2000)
Additional Comments
Genetic Interactions
Statement Reference A lilli[GD17] heterozygous background dominantly enhances the ato[1]/ato[1090] small, rough eye phenotype. A lilli[AG5] heterozygous background dominantly enhances the ato[1]/ato[1090] small, rough eye phenotype. (Distefano et al., 2012) Only 5% of abdominal segments contain a possible lch5 neuron in cato[536] ; ato[1] double homozygous embryos. (zur Lage and Jarman, 2010) amos¹ ; ato¹ double mutant embryos show a complete lack of olfactory receptor neurons in the dorsal organ ganglion. (Grillenzoni et al., 2007) Expression of Egfr::tor^act.Scer\UAS under the control of Scer\GAL4^GMR.PF in partially rescues the lack of cluster formation in the morphogenetic furrow of ato¹ eye disc clones. Expression of Egfr::tor^act.Scer\UAS under the control of Scer\GAL4^GMR.PF partially rescues the lack of cluster formation in the morphogenetic furrow of ato¹ eye disc clones. (Brown et al., 2006) ato[1]/Df(3R)p13 embryos lack C1 chordotonal organ precursor cells and have no larval oenocyte precursors. rho[Scer\UAS.cdCa]; Scer\GAL4[en-e16E] rescues the formation of larval oenocyte precursors in these embryos, but also causes additional induction and delamination of larval oenocyte precursors after the stage when this has ceased in wild-type embryos. (Brodu et al., 2004) When BacA\p35^Scer\UAS.cHa is driven by Scer\GAL4^Dll-981 in the antennal disc of ato¹/Df(3R)p13 mutants, only a partial suppression of the ectopic glial cell phenotype is seen. The fascicular organisation of the sensory afferents from the antenna is altered in mutants. (Sen et al., 2004) Expression of BacA\p35^GMR.PH increase the size of ato¹/Df(3R)p13 eyes by 25-30%. The increase in size is due to extra secondary pigment cells. (Siddall et al., 2003) Expression of sc[Scer\UAS.cCa] under the control of Scer\GAL4[7] in ato[1] flies rescues a small number of ommatidia. The eyes contain numerous interommatidial bristles in these flies. (Sun et al., 2003) When amos¹ clones are made in ato¹ animals the clone patch contains no sense organs except bristles. (zur Lage et al., 2003) The lack of oenocytes seen in ato¹/Df(3R)p13 embryos is rescued by expression of rho^{Scer\UAS.cdCa} under the control of Scer\GAL4^en-e16E. (Brodu et al., 2002) The eyes of amos^Roi-1/+ ; ato¹/ato¹ flies can reach up to 1/3 of wild-type size. The ommatidial clusters appear disorganised and contain an abnormal number of photoreceptors, whose rhabdomeres are often elongated and misshapen. R8 cells appear to be present in most if not all ommatidia. (Chanut et al., 2002) Combining Df(2L)GR4 (which removes the ASC: Achaete-scute Complex) and ato¹ causes the loss of most cells in the embryonic peripheral nervous system (PNS) - only the two dorsal located md neurons remain. (which removes the ASC: Achaete-scute Complex) (Brewster et al., 2001) ato¹/+ ; da¹⁰/+ flies have significantly fewer sensilla coeloconica on the antenna than wild-type flies. lz³; ato¹/ato¹ double mutants show a complete absence of sensilla basiconica and coeloconica, as expected from loss of lz and ato functions respectively. (Goulding et al., 2000) 1 or 2 dorsal multiple dendritic neurons remain in each hemisegment in Df(1)sc-B57; ato¹ embryos. amos^Scer\UAS.cHa rescues chordotonal neuron formation in odd segments in ato¹ embryos when expressed under the control of Scer\GAL4^h-1J3. (Huang et al., 2000) The addition of sc^t3'F.5.8 and sc^t5'eye to ato¹ flies does not restore any boss-expressing R8 photoreceptors or ommatidia. The addition of sc^t3'F.5.8 and ato^t5'eye to ato¹ flies does restore significant numbers of ommatidia and R8 receptors. The addition of ato^t3'F.5.8 and sc^t5'eye to ato¹ flies does restore significant numbers of ommatidia but does not restore R8 receptors. This leads to an unusual arrangement of photoreceptors in the ommatidia. Many of these ommatidia contain fewer than eight photoreceptors, and most of the remaining photoreceptors resemble outer photoreceptors by their positions and the large size of their rhabdomeres. The expression of sc^Scer\UAS.cCa when driven by Scer\GAL4⁷ in ato¹ flies does result in partial restoration of the compound eye. ommatidia contain only 2-5 photoreceptors, usually lacking the centrally located R8 photoreceptors with small rhabdomeres. Occasionally bristles that resemble macrochaetae are seen in various positions in the eye. (Sun et al., 2000) The addition of two copies of either or both sc^t3'F.5.8 and sc^t5'eye does not restore any boss-expressing R8 photoreceptors or ommatidia. The addition of sc^t3'F.5.8 and ato^t5'eye to ato¹ flies does restore significant numbers of ommatidia and R8 receptors, even though either transgene alone gives no rescue/suppression of this ato¹ phenotype. The addition of ato^t3'F.5.8 and sc^t5'eye to ato¹ flies does restore significant numbers of ommatidia but does not restore R8 receptors, even though either transgene alone gives no rescue/suppression of this ato¹ phenotype. This leads to an unusual arrangement of photoreceptors in the ommatidia. Many of these ommatidia contain fewer than eight photoreceptors, and most of the remaining photoreceptors resemble outer photoreceptors by their positions and the large size of their rhabdomeres. The expression of sc^Scer\UAS.cCa when driven by Scer\GAL4⁷ in ato¹ flies does result in partial restoration of the compound eye. Ommatidia contain only 2-5 photoreceptors, usually lacking the centrally located R8 photoreceptors with small rhabdomeres. Occasionally, bristles that resemble macrochaetae are seen in various positions in the eye. (Sun et al., 2000) da⁹; ato¹ double heterozygotes exhibit reduced number of antennal coeloconic sensilla. (Gupta and Rodrigues, 1997) Df(1)sc10-1 ato¹/Df(3R)p13 males do not show any decrease in the numbers of sensilla basiconica and trichodea. (zur Lage et al., 1997)
Xenogenetic Interactions
Statement Reference Expression of Bmor\ato[Scer\UAS.cYb] under the control of Scer\GAL4[ato.eye] suppresses the loss of eye phenotype found in ato[1]/Df(3R)p13 flies. However, ordered ommatidial packing is not restored as a number of supernumerary R8 cells are still present. Expression of ato::Bmor\ato[DmBm.Scer\UAS] under the control of Scer\GAL4[ato.eye] in ato[1]/Df(3R)p13 results in mild suppression of the eye degeneration phenotype (less than upon expression of ato::Bmor\ato[BmDm.Scer\UAS] or ato, Bmor\ato single mutants). Expression of ato::Bmor\ato[DmBm.Scer\UAS] under the control of Scer\GAL4[ato.eye] in ato[1]/Df(3R)p13 results in mild suppression of the eye degeneration phenotype (less than upon expression of ato, Bmor\ato single mutants). (Yu et al., 2012) Mmus\Neurog1^Scer\UAS.cQa; Scer\GAL4⁷ does not rescue eye disc phenotypes in ato¹ homozygous animals. (Quan et al., 2004) Expression of Xlae\ath5a[Scer\UAS.cSa] under the control of Scer\GAL4[7] restores the formation of R8 photoreceptor cells and numerous ommatidia in ato[1] flies. Expression of Mmus\Atoh7[Scer\UAS.cSa] under the control of Scer\GAL4[7] in ato[1] flies rescues only a small number of ommatidia (less than 25 per eye) and none of these contain photoreceptor cells with clear R8 cell morphology. Interommatidial bristles are not produced in these flies. Expression of Xlae\neuroD[Scer\UAS.cSa] under the control of Scer\GAL4[7] restores the formation of R8 photoreceptor cells and numerous ommatidia in ato[1] flies. (Sun et al., 2003) A single copy of P{UAS-Mmus\Atoh1.B} restores 3 of the 5 lateral chordotonal organs seen in wild-type, and 2 copies restores all of them, though occasionally a sixth organ is seen. (Wang et al., 2002) The chordotonal neuron phenotype is partially rescued by the expression of Mmus\Atoh1^Scer\UAS.cBa under the control of Scer\GAL4^hs.PB. (Ben-Arie et al., 2000)
Complementation & Rescue Data
Rescued by	ato¹/Df(3R)p13 is rescued by ato^Scer\UAS.cJa/Scer\GAL4^ato.3.6 (Hassan et al., 2000) ato¹/Df(3R)p13 is rescued by ato^Scer\UAS.cYb/Scer\GAL4^ato.eye (Yu et al., 2012) ato¹ is rescued by ato^t3'F.5.8/ato^t5'eye (Sun et al., 2000, Sun et al., 2000)
Partially rescued by	ato¹ is partially rescued by ato^Scer\UAS.cCa/Scer\GAL4^h-1J3 (Chien et al., 1996)
Not rescued by	ato¹/Df(3R)p13 is not rescued by Scer\GAL4^SG18.1/ato^Scer\UAS.cJa (Jhaveri and Rodrigues, 2002) ato¹ is not rescued by ato^t3'F.5.8 (Sun et al., 2000, Sun et al., 2000) ato¹ is not rescued by ato^t5'eye (Sun et al., 2000, Sun et al., 2000)
Comments	Expression of ato[Scer\UAS.cYb] under the control of Scer\GAL4[ato.eye] rescues the loss of eye phenotype found in ato[1]/Df(3R)p13 flies. However, ordered ommatidial packing is not restored as a number of supernumerary R8 cells are still present. (Yu et al., 2012) Expression of ato[Scer\UAS.cJa] under the control of Scer\GAL4[7] restores the formation of R8 photoreceptor cells and numerous ommatidia in ato[1] flies. (Sun et al., 2003) The lobular innervation pattern defect of ato¹/Df(3R)p13 flies is rescued by ato^Scer\UAS.cJa expressed under the control of Scer\GAL4^ato.3.6. (Hassan et al., 2000)
Stocks ( 1 )
Bloomington	25779 w^*; ato¹/TM6B, Tb¹
Notes on Origin
Discoverer
	Separable from: a closely linked lethal mutation. (Jhaveri and Rodrigues, 2002)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 3 )
Reported As
Symbol Synonym	ato1 (Dong et al., 2002, Ben-Arie et al., 2000) ato¹ (Sun et al., 2000, Brown et al., 2006, Sun et al., 2003, Grillenzoni et al., 2007, Wu et al., 2011, Benton et al., 2009, Brodu et al., 2004, Distefano et al., 2012, Gopfert et al., 2002, zur Lage and Jarman, 2010, Witt et al., 2010, Cachero et al., 2011, Ma and Jarman, 2011, Bardin et al., 2010, Melicharek et al., 2008, Yu et al., 2012, Plavicki et al., 2012, Senthilan et al., 2012)
Name Synonym	atonal¹ (Cosetti et al., 2008)
Secondary FlyBase IDs

References ( 67 )

Generate a list of	All references relating to this allele ( 67 )

List References by type	Research paper ( 66 ) Review ( 1 )
Recent research papers ( 7 )
	Distefano et al., 2012, Dev. Dyn. 241(3): 553--562 Drosophila lilliputian is required for proneural gene expression in retinal development. [FBrf0217520] Plavicki et al., 2012, Proc. Natl. Acad. Sci. U.S.A. 109(5): 1578--1583 Homeobox gene distal-less is required for neuronal differentiation and neurite outgrowth in the Drosophila olfactory system. [FBrf0217395] Senthilan et al., 2012, Cell 150(5): 1042--1054 Drosophila auditory organ genes and genetic hearing defects. [FBrf0219321] Yu et al., 2012, genesis 50(5): 393--403 Identification of Bombyx atonal and functional comparison with the Drosophila atonal proneural factor in the developing fly eye. [FBrf0218282] Cachero et al., 2011, PLoS Biol. 9(1): e1000568 The gene regulatory cascade linking proneural specification with differentiation in Drosophila sensory neurons. [FBrf0212891] Ma and Jarman, 2011, J. Cell Sci. 124(15): 2622--2630 Dilatory is a Drosophila protein related to AZI1 (CEP131) that is located at the ciliary base and required for cilium formation. [FBrf0214370] Wu et al., 2011, Neuron 70(2): 281--298 A combinatorial semaphorin code instructs the initial steps of sensory circuit assembly in the Drosophila CNS. [FBrf0213571] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2011 Show reviews ...

Allele Dmel\ato1

Allele Dmel\ato¹